Estimating sensory delays to primate M1: a comparison of peri-stimulus time histograms and coherence phase-frequency regression Average PSTH Boubker Zaaimi, Demetris S. Soteropoulos, Elizabeth R. Williams & Stuart N. Baker Institute of Neuroscience, Newcastle University, UK. E-mail: [email protected] Introduction 277.3/KK29 Experimental Results 50 0 -20 50 100 150 200 0 Time after stimulus (ms) Coherence 0.06 50 100 Time after stimulus (ms) Coherence 0.5 0.04 0.02 0.4 0.3 0.2 60 80 100 0.02 0 0 10 20 30 Frequency (Hz) 40 50 0 40 60 Frequency (Hz) Coherence Phase 80 0 100 20 40 60 80 Frequency (Hz) 100 Coherence Phase 3p Phase (rad) 2p p 2p p 40 60 Frequency (Hz) 80 100 40 60 80 20 30 Frequency (Hz) 40 50 0 20 40 Time(ms) 60 80 100 Probability per bin Results from simulations with the same response onset latency, but with increasing response duration. The latency estimated from the coherence phase-freqeuncy regression (arrowheads) was always close to the middle of the response. 0.03 0.02 0 0.01 0 Spike train simulated as 2 inhomogeneous gamma process (order parameter = 4) -20 0 20 40 60 80 100 Spike Response 0.01 0.018 Fraction of intervals 0.006 0.004 0.002 0.014 50 100 150 Inter-stimulus interval (ms) 200 0.03 0.02 40 Time(ms) 60 40 Time(ms) 60 80 100 0.03 0.02 0.01 0 -20 0 20 80 100 Physiological responses often include an initial brief facilitation, followed by a long lasting, but weak, suppression (e.g. the experimentally recorded example cell A). In a simulation mimicking this, the coherence delay estimate was substantially larger than the onset latency, as it was affected by the suppression. Using coherence phase-frequency relationships to estimate delay measures the average delay of the whole response, not just the onset latency. This must be remembered when comparing coherence delays with measurements in the literature, which are usually based on the earliest response to a stimulus. In some circumstances, coherence delay estimates may be more functionally relevant than those based on onset latency. 0.01 0.01 0 -20 0 20 40 0.006 60 80 100 References Time(ms) 0.002 0 0.04 Probability per bin Spike train 20 Conclusion Time(ms) 0.008 100 0.02 0.01 Probability changes sum to give instantaneous spiking probability Stimulus 80 0.04 0.01 0.04 100ms 60 0.02 0.04 Fixed change in spiking probability follows each stimulus 0 40 Time(ms) When the response included both a facilitation and suppression, coherence delay estimate was close to the centroid of the absolute response profile (i.e. Ignoring its sign). -20 Simulation Methods 0.02 20 0.03 0 -20 Firing Probability per 1ms bin 0 0.04 100 Frequency (Hz) Delays estimated from coherence phase-frequency slopes are substantially higher than from PSTH onset latency: in the examples above 18 vs 10 ms for the median nerve stimulation, and 17 vs 10 ms for the digital nerve stimulation. We investigated this further using simple computer simulations. 0 -20 0 20 0 Poisson stimulus train 10 0 0.04 Probability per bin 20 0 0 0.02 0.03 -p 0 -p -2p -3p 0 -p -2p -3p 0 Phase p 0.03 0.01 Probability per bin 20 Phase (rad) 0 Coherence was greater at low frequencies, reflecting the low-pass filtering consequences of the response duration. For a more complex response profile, the coherence delay estimate was close to the centroid (centre of mass) of the response. 0.04 Phases differed by p radians between facilitation and suppression. Regression slopes implied a delay of 14.5ms (arrowheads on PSTH). 0.04 0.1 0 Fraction of intervals 40 Coherence Coherence 0 1s 20 Time(ms) Poisson stimulus train 1 mV 0 0 0 3p Microelectrode penetrations into primary motor cortex of a macaque monkey, sedated using ketamine/medetomidine. Single unit activity isolated. Electrical stimulation of median nerve at the arm (biphasic stimuli, 0.2ms per pulse, intensity at motor threshold), or digital nerve of index finger (10mA) 1 Stimuli delivered as Poisson stimulus trains , mean rate 10Hz. 0.002 20 Phase (rad) ! Counts Counts 40 Methods ! 100 60 In this poster, we examine this assumption. ! Peri-stimulus Time Histogram Examples of delay estimated from coherence phase for more complex response profiles. Each plot shows the PSTH, and the coherence delay estimate (arrowhead). Probability per bin 80 Coherence It is commonly assumed that coherence phase analysis will yield similar delay estimates to response latencies, measured using timedomain methods such as the peri-stimulus time histogram (PSTH). B Peri-stimulus Time Histogram Coherence For a system with a fixed delay, phase (q) is linearly related to frequency (f) with a slope related to the delay (t): q(f) = 2ptf Spike probability per bin Coherence is a measure of correlation in the frequency domain between two signals. Coherence phase indicates the average phase difference between the signals at a given frequency. Digital nerve stimulation Median nerve stimulation A Example simulation results where cell responded with a brief facilitation (red) or suppression (blue) of its firing (onset latency 9.5ms, duration 10ms) Probability per bin PSTH 0.004 Coherence analysis is widely used to investigate oscillatory phenomena in the nervous system. 0 0 50 100 150 Inter-spike interval (ms) 200 1. Miller S, Clark J, Eyre JA, Kelly S, Lim E, McClelland VM, Mc Donough S & Metcalfe AV. (2001). Comparison of spinal myotatic reflexes in human adults investigated with cross-correlation and signal averaging methods. Brain Res 899, 4765. 2. Baker SN & Gerstein GL. (2000). Improvements to the sensitivity of gravitational clustering for multiple neuron recordings. Neural Comput 12, 2597-2620.
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